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Morphological Character Mapping on a Molecular Phylogeny Using Pollen Variation in the Cryptanthinae (Boraginaceae)

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Morphological Character Mapping on a Molecular Phylogeny Using Pollen Variation in the Cryptanthinae (Boraginaceae) Morphological Character Mapping on a Molecular Phylogeny Using Pollen Variation in the Cryptanthinae (Boraginaceae) by Rachel Spaeth A thesis submitted to Sonoma State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE in Biology ______________________ Dr. Richard Whitkus, Chair ______________________ Dr. Michelle Goman ______________________ Dr. Murali Pillai ______________________ Date Copyright 2014 By Rachel Spaeth ii Authorization for Reproduction of Master’s Thesis Permission to reproduce this thesis in part or entirety must be obtained from me. DATE:________________________ ______________________________ Signature ______________________________ Street Address ______________________________ City, State, Zip iii Morphological Character Mapping on a Molecular Phylogeny: Using Pollen Variation in the Cryptanthinae (Boraginaceae) Thesis by Rachel Spaeth ABSTRACT Phylogenetic classification at the species level in the Boraginaceae is notoriously difficult when relying solely on morphological data. Studies are currently in progress to generate a well-supported phylogenetic tree of this family using molecular data. A molecular based phylogeny may reveal the characters that evolved slowly enough to have the same state in closely related taxa found in some key palynological traits used in previous classifications. Pollen attributes were collected on seventy four species across six genera in the subtribe Cryptanthinae using Scanning Electron Microscopy (SEM). The pollen data exhibit features which are taxonomically informative including shape, aperture type, sculpturing, and size. Cryptanthinae pollen encompasses three of the nine Erdtman (1966) shape categories, and seven of the eleven Faegri and Iversen (1975) sub- shape categories. Their aperture types include heterocolpate, zonoporate, and zonocolpate forms. They are sculpted with fossulate, foveolate, echinate, reticulate, and gemmate clavate surfaces. They range in size from 4.85μm long and 1.92μm wide to 40.85μm long and 25.60μm wide. Some of them have a transverse groove and others do not. The same is true for the presence or absence of polar apertures. These characteristics were mapped on a molecular phylogeny to observe evolutionary trends. Biogeographic data such as habitat moisture, range of distribution, flowering period, and style type were also mapped on the molecular phylogeny to uncover selection pressures responsible for the high level of morphological diversity in this subfamily. This analysis revealed habitat moisture level as one of the driving forces behind pollen subshape diversity in the Cryptanthinae. Chair: _________________________ Signature MS Program: Biology iv Sonoma State University Date: ___________ Acknowledgement Funding provided by the California Rare Fruit Growers Association Redwood Empire Chapter and the California Native Plant Society Milo Baker Chapter. Special thanks to Dr. Richard Whitkus, Matthew Guilliams, Steve Anderson, Dr. Murali Pillai, Dr. Michelle Goman, Daniel Streeter, The William M. Keck Microanalysis Laboratory, the Northcoast Herbarium, the University of California at Berkeley / Jepson Herbarium, Universidad de Conceptión Herbario, and Missouri Botanical Garden Herbarium. v Table of Contents Introduction…………………………………..……………………..…...………………... 1 Research Questions………………………………………………..…..………………...... 9 Methods…………………………………………………………………..………………..9 Results……………………………………………………….…………………………... 15 Morphological Variation………………………………………....……....................... 15 Pollen Size………………………………………………………………………... 15 Pollen Shape…………………………………………………………………….... 17 Pollen Subshape……………………………………….…………………………..18 Pollen Apertures……………………………………….….……………………… 20 Pollen Sculpturing…………………………………………………....................... 21 Phylogenetic Trait Distribution………………………………….……........................ 22 Pollen Size………………………………………………...……………………… 24 Pollen Shape……………………………………………….................................... 24 Pollen Subshape………………………………………….......................................26 Pollen Apertures………………………………….……….……………………….28 Pollen Sculpturing………………………………………....……………………... 30 Habitat, Biogeography, and Breeding System………………….……......................... 32 Habitat Moisture…………………………………………………..……………… 32 Range…………………………………....…………….....…………...................... 34 Flowering Period…………………………………………..................................... 36 Breeding System……………………………………………………...................... 36 Biogeographic Correlation with Morphological Features……………………………. 36 vi Discussion……………………………………………………………...……………….. .38 Cryptanthinae and Geologic Time…………………………………………………… 38 Noteworthy Cryptanthinae Clades…………………………………………………… 39 Conclusion………………………………………………….…………...………………. 44 Literature Cited………………………………………………………….......................... 45 Appendix 1………………………………………………….…………..……………….. 51 Appendix 2…………………………………………………………..….……………….. 54 Appendix 3…………………………………………………………...………………….. 67 Appendix 4……………………………………………………..…….………………….. 80 Appendix 5………………………………………………………………......................... 81 Appendix 6……………………………………………….………………........................ 81 Appendix 7………………………………………………………………………………. 82 Appendix 8………………………………………………………………………………. 82 Appendix 9………………………………………………………………………………. 83 Appendix 10……………………………………………………………………………... 83 Appendix 11……………………………………………………………………………... 84 Appendix 12……………………………………………………………………………... 86 vii List of Tables Table 1. Pollen size variation observed across genera in the subtribe Cryptanthinae (Boraginaceae). Polar (P) and equatorial (E) lengths were assessed as shown in Fig. 4. Polar to equatorial ratio (P:E) ranges represent the minimum and maximum values found among species in each genus. The Myosotis outgroup species are included as well. Page 16. Table 2. Cryptanthinae taxa sampled with Herbarium and accession information. Abbreviations are as follows: NCC - North Coast California, UC/JEPS – University of California at Berkeley/Jepson, UAZ – University of Arizona, CONC – Universidad de Concepción Peru, MO – Missouri Botanical Gardens. Page 51-53. Table 3. Pollen shape and subshape distribution among 74 taxa in the Cryptanthinae. The taxa are organized with regard to Erdtman (1966) shape as well as Faegri and Iversen (1975) subshape. The three different shapes are based on P:E ratios. The subshapes are based on the measurements taken in Fig. 4. The Myosotis outgroup species are included as well. Page 80. Table 4. Pollen aperture types found across 74 taxa of the Cryptanthinae. The Myosotis outgroup species are included as well. Page 81. Table 5. Presence or absence of transverse grooves in 74 taxa of Cryptanthinae pollen. The Myosotis outgroup species are included as well. Page 81. Table 6. Presence or absence of polar pseudo-apertures across 74 taxa of Cryptanthinae pollen. The Myosotis outgroup species are included as well. Page 82. Table 7. Sculpturing types for pollen of 74 taxa in the Cryptanthinae. The Myosotis outgroup species are included as well. Page 82. Table 8. Habitat moisture level for 74 Cryptanthinae taxa and two Myosotis outgroup taxa. Page 83. Table 9. Flowering period in selected Cryptanthinae taxa. Page 83. Table 10. Table 10. Average polar and equatorial diameter, P:E ratio, standard deviation of P and E. A standard test of normality for both P and E was performed in excel and all were normal with a significance of p<.05. Page 84-85. viii List of Figures Figure 1. General flowering characteristics of members of the Boraginoideae including cymose-based terminal inflorescence, radial corolla, superior ovary, epipetalous stamens, and nutlets (Watson and Dallwitz 2013). Page 2. Figure 2. Current placement of the Boraginaceae in the Boraginales clade (APGIII 2014). Page 3. Figure 3. Recent phylogeny of the Cryptanthinae. ‘Maximum likelihood (ML) analysis with bootstrap values (≥ 70%) indicated at nodes’ (Hasenstab-Lehman and Simpson 2012). Note the polyphyletic dispersion of Plagiobothrys and Cryptantha. Page 5. Figure 4. Pollen measurements to determine sub-shape: polar length (P), equatorial width (E), polar width (A), distance from outer polar width to maximum width (B), length from maximum width to pole (C), distance between outside maximum width (D), and maximum width (W). These line drawings show the most commonly observed sub- shapes, but are not representative of all the variation present. Page 11. Figure 5. Faegri and Iversen (1975) sub-shape classes with grayed-in sub-shapes representing those found in the Cryptanthinae subtribe. Page 12. Figure 6. Molecular phylogeny for the Cryptanthinae subtribe (Guilliams 2013) used throughout this study with era, period, and epoch information added to the figure (Polly et al. 2011). The scale axis at the bottom of the figure is in millions of years. Genera abbreviations: Pl. – Plagiobothrys; Pe. – Pectocarya; H. – Harpagonella; C. – Cryptantha; A. – Amsinckia; M. – Myosotis are used throughout this report. Page 13. Figure 7. Pollen size variability across genera in the Cryptanthinae ranges from 40.85µm-4.85µm in polar length. Maximum and minimum sizes are represented for each genus. Species are as follows: A. A. vernicosa, B. A. menziesii, C. M. discolor, D. Pl. albiflorus, E. Pe. pusilla, F. C. confertiflora, G. M. laxa, H. Pe. setosa, I. C. muricata, J. Pl. humilis. See Table 1 for sizes. Page 17. Figure 8. The range of Erdtman (1966) pollen shapes in the Cryptanthinae for Plagiobothrys, Pectocarya, and Cryptantha based on P:E ratio. Scale bars for each image are included. Species are as follows: A. Pl. uncinatus,
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